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, OCTOBER 2014, 65, 5, 375—386 doi: 10.2478/geoca-2014-0026
Introduction
Post-sedimentional erosional processes and Cretaceous sedi-
mentation removed or buried the Upper Jurassic deposits of
the Bohemian Massif. The depositional conditions, ecosys-
tem composition, and biogeographical affinity of this region
thus remain weakly known. However, a few outcrops captur-
ing the Upper Jurassic deposits are still exposed or fossil
specimens from older outcrops are preserved in museum col-
lections. Such findings allow us to fill the gap in distribution
patterns of Upper Jurassic marine organisms. Although the
first Upper Jurassic ammonites were found in the Bohemian
Massif in the 1860s (Lenz 1870; Bruder 1881, 1882, 1885,
1886), the regional composition and temporal distribution of
ammonite assemblages as well as their compositional rela-
tionships with adjacent geographical regions were poorly ex-
plored. In this paper, we focus on systematic revision of
museum collections of two ammonite families of high bio-
stratigraphic and paleobiogeographic significance (Aula-
costephanidae and Cardioceratidae), and compare them with
co-eval Tethyan and Boreal ammonite assemblages from
other regions. We note that the majority of specimens in the
ammonite assemblages in the Bohemian Massif, represented
by Perisphinctidae, Ataxioceratidae, and Oppeliidae, will be
described in a separate study.
The systematics and paleobiogeographic significance of
Sub-Boreal and Boreal ammonites (Aulacostephanidae
and Cardioceratidae) from the Upper Jurassic of the
Bohemian Massif
JAN HRBEK
Institute of Geology and Paleontology, Faculty of Science, Charles University in Prague, Albertov 6, 128 43 Prague 2, Czech Republic;
hrbek.honza@seznam.cz
(Manuscript received March 1, 2014; accepted in revised form October 7, 2014)
Abstract: Upper Jurassic marine deposits are either rarely preserved due to erosion or buried under younger sediments
in the Bohemian Massif. However, fossil assemblages from a few successions exposed in northern Bohemia and Saxony
and preserved in museum collections document the regional composition of macro-invertebrate assemblages and thus
provide unique insights into broad-scale distribution and migration pathways of ammonites during the Late Jurassic. In
this paper, we focus on the systematic revision of ammonites from the Upper Oxfordian and Lower Kimmeridgian
deposits of northern Bohemia and Saxony. The ammonites belong to two families (Aulacostephanidae and Cardioceratidae)
of high paleobiogeographic and stratigraphic significance. Six genera belong to the family Aulacostephanidae (Prorasenia,
Rasenia, Eurasenia, Rasenioides, Aulacostephanus, Aulacostephanoides) and one genus belongs to the family
Cardioceratidae (Amoeboceras). They show that the Upper Jurassic deposits of the northern Bohemian Massif belong to
the Upper Oxfordian and Lower Kimmeridgian and paleobiogeographically correspond to the German-Polish ammonite
branch with the geographical extent from the Polish Jura Chain to the Swabian and Franconian Alb. Therefore, the
occurrences of ammonites described here imply that migration pathway connecting the Polish Jura Chain with habitats
in southern Germany was located during the Late Oxfordian and Early Kimmeridgian in the Bohemian Massif.
Key words: Oxfordian, Kimmeridgian, Czech Republic, biostratigraphy, paleobiogeography, ammonites,
Aulacostephanidae, Cardioceratidae.
Fig. 1. Schematic map with surface outcrops of Jurassic rocks and
Lusatian fault course in northern Bohemia (modified according to
Kopecký et al. 1963).
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In the Late Jurassic, the Sub-Mediterranean Realm was lo-
cated along the northern margin of Tethys (namely the Euro-
pean part in the vicinity of the Mediterranean Sea – southern
Spain, southern France, peri-Carpathian area and Balkan
area in the east). The Sub-Boreal Realm ranged from Scot-
land, southern England, northern France, northern Germany,
northern Poland up to the southern part of the Russian plat-
form and Caspian area. The Boreal Realm was located north
of the Sub-Boreal Realm. It was characterized by high-lati-
tude, cold-water faunas occurring in the Canadian Arctic,
Alaska, Greenland, Spitsbergen, the Barents Sea, and Siberian
area (Callomon 2003).
Aulacostephanids are typical of the Sub-Boreal Realm and
are used for zonal and subzonal subdivisions of the Sub-Bo-
real Upper Oxfordian and Kimmeridgian (Birkelund & Cal-
lomon 1985; Krymholts et al. 1988; Wright 2010). However,
aulacostephanids also occur in the Sub-Mediterranean and
Boreal regions. In contrast, Cardioceratids are typical of the
Boreal Realm at high latitudes, and occur less frequently in
Sub-Boreal and Sub-Mediterranean realms. The proposed
global stratotype for the Oxfordian/Kimmeridgian (O/K)
boundary, located at Staffin Bay, Isle of Skye in Scotland, is
characterized by numerous aulacostephanid occurrences
below and above this boundary (Matyja et al. 2006;
Wierzbowski et al. 2006). This boundary is characterized by
the replacement of the aulacostephanid Ringsteadia and
Microbiplices by Pictonia and Prorasenia, which coincides
with the first appearance of the Boreal species Amoeboceras
(Plasmatites) (Wierzbowski et al. 2006; Glowniak et al.
2010; Wierzbowski 2010).
Materials and methods
This study is based on specimens known from the collec-
tions of the National Museum, Prague (NM-N), the Institute
of Geology and Paleontology, Faculty of Science, Charles
University, Prague (CIGP) and the State Museum of Geology
and Paleontology, Dresden (SMGPD). This paper includes
descriptions of 26 specimens belonging to Aulacoste-
phanidae and Cardioceratidae, mostly coming from the
Czech Republic, partly including specimens recorded in Ger-
many. The ammonites described here were collected in the
vicinity of Šluknov, Krásná Lípa, Doubice, Brtníky and
Kyjov, in the former Šternberk quarry (northern Bohemia)
and in Hohnstein, Saxony (Germany) (Fig. 1).
Ammonite preservation does not allow the determination of
morphological characters in the majority of specimens. Su-
tures and other surface structures are not preserved, and deter-
mination of the body chamber and phragmocone part is
hampered by poor preservation of internal moulds. Determi-
nation to a genus level is not possible in some specimens owing
to strong shell deformation. However, the morphological char-
acters of shells are sufficiently preserved in some specimens
and allow comparison with aulacostephanids from Europe.
The following abbreviations are used in the ‘Measure-
ments’ chapter:
D (max)
maximum diameter preserved
D (phg-ad)
diameter of fully grown individual
D (phg)
diameter of final whorl (incomplete conch)
Wh/D
ratio of height and diameter
Wt/D
ratio of thickness and diameter
U/D
ratio of umbilicus and diameter.
Geological settings
Jurassic outcrops in northern Bohemia are exposed only at
several places as relicts of the epi-continental sea. These out-
crops are associated with the Lusatian Fault that is responsi-
ble for overturning of successions and probably contributed
to tectonic alteration and dolomitization (Eliáš 1981). Dur-
Fig. 2. The original lithostratigraphic scheme with lithotypes col-
lected near Krásná Lípa town. Two massive limestone beds (Bimam-
matum and Platynota Zones) represent the most significant parts with
high density of fauna (modified according to Bruder 1882, 1886).
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ing the younger phase of the Saxonic tectonic event, the
re-activation of the Lusatian tectonic zone pulled the Cado-
mian, Paleozoic and Jurassic sediments over the Cretaceous
sediments.
Carboniferous and Permian sediments form the bedrock of
the Middle Jurassic Brtníky Formation (Suk et al. 1984),
which reaches up to 20 meters in thickness and is formed
predominantly by sandy, fossil-poor clastics. The onset of
sedimentation probably corresponds to the Callovian (Bruder
1881, 1882; Kopecký et al. 1963; Eliáš 1981; Chlupáč et al.
2002). The Doubice Formation overlies sandstones of the
Brtníky Formation and exceeds 100 m in thickness (Kopecký
et al. 1963; Eliáš 1981; Suk et al. 1984). It is predominantly
formed by micritic limestones that were deposited in a deep-
shelf marine environment. The lithological composition is
variable, formed by interbeds of marly limestones, cherts,
marls, and sandy rocks – see origin profile (Fig. 2). Carbon-
ates are weakly to strongly dolomitized (due to influence of
the Lusatian tectonic zone and the Tertiary volcanism). The
Doubice Formation contains diverse fossil assemblages with
ammonites, belemnites, brachiopods, bivalves, sponges, for-
aminifera, echinoderms, bryozoans, annelids and fish (Bruder
1881, 1882, 1885, 1886; Eliáš 1981). A relatively high abun-
dance of sponges (also documented in limestone blocks pre-
served in the creek below the “Peškova stráň” locality) can
imply that some assemblages correspond to sponge megafa-
cies. However, a sedimentological analysis of Upper Jurassic
facies from northern Bohemia is not possible at present due
to the lack of surface outcrops.
Near Doubice, outcrops expose overturned successions
that are located close to the Lusatian fault. Pešek’s Hillside
near Kyjov is a small-scale oucrop formed by sandy lime-
stones, marly carbonates and micritic limestones. Sparitic
limestones contain recrystallized bioclasts, mainly sponges
and bivalves. Most bioclasts occur in micritic limestones
(sponge spicules, brachiopods). Matrix and bioclasts (mainly
bryozoans, sponges and bivalves) are characterized by exten-
sive recrystallization. Some thin-sections show shear-like
structures, possibly reflecting partial redeposition. Peloids
and pellets-like structures are present (Fig. 3).
The location of coastal edge of this epicontinental sea is
unknown; possibly extensive Jurassic sediments were eroded
during the Early Cretaceous and at the beginning of the Late
Cretaceous. However, the existence of a connection between
Fig. 3. Thin sections obtained from the Late Jurassic lithotypes collected from the Peškova stráň hillside near Kyjov. A, B, C – Micritic
limestones: A, B – Predominantly almost oval-shaped peloid-like objects occur very abundantly with recrystallized organic relicts (sponges,
brachiopods); C – Strongly recrystallized bioclasts in a homogeneous matrix. D – Calcareous-clayey siltstone. Sparitic siltstone with
fragments of quartz suggests the possible contribution of terrestrial material. Recrystallized faunal relicts (bivalves, gastropods, corals,
sponges) occur at a lower density in comparison with limestones. Jurassic lithotypes are characterized by similar development, but differ in
degree of recrystallization.
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the northern Boreal Realm in the NW and the Tethyan
Realm in the SE is suggested by the composition of ammo-
nite assemblages. The marine ways, which led probably
through the Paris Basin and Germany connecting the Tethyan
Realm with the Boreal Realm (Fig. 4), were first opened dur-
ing the Callovian (Page et al. 2009). During the Late Callov-
ian, the Boreal sea penetrated into Bohemia from the north
via the Labe zone and formed the Moravian (Saxonian)
strait, which represents one of the most important migration
routes in Europe during the Late Jurassic (Eliáš 1981).
Systematic paleontology
Order: Ammonitida Zittel, 1885
Superfamily: Perisphinctoidea Steinmann, 1890
Family: Aulacostephanidae Spath, 1924
Genus: Microbiplices Arkell, 1936
Microbiplices sp.
Fig. 5A
M a t e r i a l : NM-N55; from Šternberk quarry near Brtníky.
M e a s u r e m e n t s : NM-N55: D (phg): 28 mm; Wh/D: 0.30;
Wt/D: 0.35; U/D: 0.40.
D e s c r i p t i o n : Carbonate internal mould with preserved
inner whorls. The umbilical part and lateral sides are not pre-
served. Evolute type of shell and regular rib bifurcation is
typical. Umbilicus is wide. The whorl section is round to
oval. Primaries have an almost radial direction. Secondary
ribs cross the ventral margin fluently without fading.
R e m a r k s : Carbonate internal mould, gently deformed.
Character of ribbing and shell coiling suggests some affinity
to Microbiplices microbiplex figured by Matyja et al. 2006,
fig. 4a and M. microbiplex figured by Schairer 2003 (pl. 2,
fig. 7, 8).
Genus: Prorasenia Schindewolf, 1925
Prorasenia sp.
Fig. 5B—E
M a t e r i a l: NM-N62/1, (fig. 5B); Šternberk near Brtníky.
NM-N183 (fig. 5C); Kyjov near Krásná Lípa. NM-N56
(fig. 5D); Šternberk quarry near Krásná Lípa. NM-N62/2
(fig. 5E); Šternberk quarry near the Brtníky village.
M e a s u r e m e n t s : NM-N62/1 (No. 62): D (max): 22 mm.
NM-N183; D (max): 36 mm. NM-N56; D (max): 27 mm;
Wh/D: 0.23; Wt/D: 0.23, not measured. NM-N62/2; D (max):
12 mm.
D e s c r i p t i o n : Evolute types of shell with low density of
ribbing, only the specimen 5B shows a somewhat higher
density of ribbing. Primaries regularly bifurcate, between
them some ribs trifurcate, which is typical of this genus. It is
visible in the specimens 5C and 5E. Ribs are typically sharp
and they have almost radial direction.
R e m a r k s: NM-N62/1: This specimen suggests a some-
what more prorsiradiate ribs than other figured specimens. It is
similar to Rasenia cf. hardyi figured by Schweigert & Cal-
lomon, 1997, pl. 7, fig. 10 and Rasenia (Prorasenia) n. sp.
figured by Geyer, 1961, pl. 1, fig. 8. NM-N183: Low density
of ribbing suggests the similarity with the Prorasenia sp. fig-
Fig. 4. Map showing paleogeographical situation in Central Europe during the Late Jurassic (modified according to Matyja & Wierzbowski
1995).
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Fig. 5. Ammonites of the Microbiplices, Prorasenia, Rasenia, Aulacostephanoides and Aulacostephanus group. A – Microbiplices sp.,
NM-N55, Kyjov. B—E – Prorasenia sp.: B – NM-N62/1, Šternberk quarry; C – NM-N183, Kyjov; D – NM-N56; E – NM-N62/2, both
from Šternberk quarry. F—H – Prorasenia cf. bathyschista Koerner: F – adult specimen; F1 – view from the right; F2 – ventral view,
NM-N173, Kyjov; G1 – view from the right, G2 – view from the left, CIGP 12, Šternberk quarry; H – SMGPD CsJ 45, Doubice. I – Pro-
rasenia crenata Quenstedt, CIGP 10506, Šternberk quarry. J – Rasenia sp. – NM-N179, Kyjov. K – Aulacostephanoides sp.: K1 – view
from the left, K2 – view from the right, K3 – ventral view, SMGPD SaJ 3, Hohnstein quarry (Germany). L – Aulacostephanus sp.:
L1 – view from the right, L2 – view from the left, SMGPD SaJ 15, Hohnstein quarry (Germany).
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ured by Schairer, 2003 (pl. 2, fig. 12). Dichotomical and tri-
chotomical branching of ribs is present. Both samples have
similar shell size. NM-N56: Specimen is deformed and in-
complete. It shows some similarity to Prorasenia crenata fig-
ured by Wierzbowski, 2010 (pl. 10, fig. 3, 4). NM-N62/2: One
trichotomically branched rib suggests a similarity with genus
Prorasenia. The density of ribs is not high. Prorasenia cf.
hardyi figured by Schweigert & Callomon, 1997 (pl. 7,
fig. 10) has somewhat stronger ribs than the specimen 5E.
Prorasenia cf. bathyschista Koerner, 1963
Fig. 5F—H
M a t e r i a l : NM-N173; (fig. 5F) Kyjov near Krásná Lípa.
CIGP 12; (fig. 5G) former Šternberk quarry. SMGPD CsJ 45
(fig. 5H); Doubice village near Krásná Lípa.
M e a s u r e m e n t s : NM-N173; D (phg-ad): 47 mm; Wh/D:
0.35; Wt/D: 0.32; U/D: 0.45. CIGP 12; D (max): 33 mm;
Wh/D: 0.30; Wt/D: 0.37, not measured. SMGPD CsJ 45;
D (phg): 30 mm; Wh/D: 0.36; Wt/D: 0.37; U/D: 0.42.
D e s c r i p t i o n : Evolute shells with a round whorl cross-
section. Primary ribs are slightly prorsiradiate, secondaries
bend gently backward and form S-shaped deflection.
Re m a r k s : NM-N173: Prorasenia cf. bathyschista closely
resembles the species figured by Matyja & Wierzbowski,
1998 (pl. 2, fig. 4, 5).
Prorasenia cf. crenata Quenstedt, 1887
Fig. 5I
M a t e r i a l : CIGP 10506, from Šternberk quarry.
M e a s u r e m e n t s : D (max): 35 mm; Wh/D: 0.26; Wt/D:
0.28; U/D: 0.37.
D e s c r i p t i o n : Evolute coiling with the umbilical part
and a round whorl cross-section. Primary ribs are gently
prorsiradiate, sharp, and split into two secondaries, which
have backward direction.
R e m a r k s : The specimen is heavily deformed and the
shell is incomplete. The ribbing suggests that it can belong to
Prorasenia. The depicted specimen is of small size (less than
50 mm in diameter), similarly as Prorasenia crenata figured
by Wierzbowski, 2010 (pl. 10, fig. 1—5). The character of rib-
bing is very similar in both specimens. Ribs are weaker then in
P. hardyi figured by Schweigert & Callomon, 1997 (pl. 7,
fig. 10), but the rib shape is similar in both specimens.
O c c u r r e n c e : Prorasenia crenata occurs in the upper
part of the Bimammatum Zone (the Bimammatum and the
Hauffianum Subzones) and the lower part of the Planula Zone.
Genus: Rasenia Salfeld, 1913
Rasenia sp.
Fig. 5J
M a t e r i a l : NM-N179; Kyjov near Krásná Lípa.
M e a s u r e m e n t s : NM-N179; D (phg): 38 mm; Wh/D:
0.26; Wt/D: 0.26; U/D: 0.47.
D e s c r i p t i o n : The character of inner whorls is reduced,
ribbing is not apparent. The whorl cross-section is square-
shaped up to oval with flat lateral sides. Umbilical wall is
very low or absent. Primary ribs have radial direction. On the
very flat ventral side, the secondaries appear with the same
direction as primaries.
R e m a r k s : Gently deformed, internal mould is cracked.
Inner whorl is poorly preserved and does not allow detailed
determination.
Genus: Eurasenia Geyer, 1961
Eurasenia sp.
Fig. 6A—C
M a t e r i a l : NM-N184, (fig. 6A) found in Šternberk quarry
near Krásná Lípa. CIGP 16305, (fig. 6B) found in Hohnstein
quarry (Germany). SMGPD SaJ 7, (fig. 6C) found in Hohnstein
quarry, Germany.
M e a s u r e m e n t s : NM-N184; D (max): 175 mm. CIGP
16305; D (max): 54 mm. SMGPD SaJ 7; D (phg): 28 mm;
Wh/D: 0.31; Wt/D: 0.30; U/D: 0.37.
D e s c r i p t i o n: Very strong, massive and short primary
ribs suggest some affinity to Eurasenia. Our specimens dif-
fer in the direction of ribbing. The specimen 2A has gently
prorsiradiate ribs, 2B has almost radial ribs, and 2C shows
slightly rursiradiate type of ribbing. Umbilicus is narrow
with involute coiling of whorl. The whorl cross-section
seems to be oval to square-shaped.
R e m a r k s : NM-N184: The specimen is similar to Rasenia
(Eurasenia) pendula figured by Geyer, 1961 (pl. 9, fig. 7).
R. (E.) frischlini by Geyer, 1961 (pl. 18, fig. 2) is also simi-
lar to the specimen figured here. CIGP 16305: Rasenia
(Eurasenia) trimera figured by Geyer, 1961 (pl. 9, fig. 6)
seems to be very similar to the sample figured here, mainly
by massive primary ribs and splitting into secondaries.
SMGPD SaJ 7: Very short and massive primary ribs and their
splitting is strikingly similar to Rasenia (Eurasenia) balteata
figured by Geyer, 1961 (pl. 18, fig. 1), but differs by having
more rursiradiate ribs than the specimen figured here.
Genus: Rasenioides Schindewolf, 1925
Rasenioides sp.
Fig. 6D—H
M a t e r i a l : SMGPD SaJ 12 (fig. 6D), found in Hohnstein
quarry (Germany). SMGPD SaJ 24 (fig. 6E), found in
Hohnstein quarry, (Germany). NM-N96 (fig. 6F), Šternberk
quarry near Brtníky village. SMGPD SaJ 25 (fig. 6G), found
in Hohnstein quarry, (Germany). NM-N60, (fig. 6H); Kyjov
near Krásná Lípa town.
Measurements: SMGPD SaJ 12; D (phg): 29 mm; Wh/D:
0.35; Wt/D: 0.29; U/D: 0.42. SMGPD SaJ 24; D (phg):
22 mm, Wh/D: 0.35; Wt/D: 0.32; U/D: 0.28. NM-N96;
D (phg): 41 mm; Wh/D: 0.32; Wt/D: 0.40 U/D: ?0.45.
SMGPD SaJ 25; D (phg): 30 mm; Wh/D: 0.38; Wt/D: 0.20;
U/D: 0.31. NM-N60: D (phg): 40 mm; Wh/D: 0.39; Wt/D:
0.28; U/D: 0.33.
D e s c r i p t i o n : Rasenioides is characterized by involute
type of shell with short and strong primary ribs and by high
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Fig. 6. Ammonites of the Eurasenia, Rasenioides, Vielunia and Amoeboceras group. A—C – Eurasenia sp.: A – NM-N184, view from the
right, Šternberk quarry; B – CIGP 16305, fragment of the whorl; C – SMGPD SaJ 7, both from Hohnstein quarry (Germany). D—H – Ra-
senioides sp.: D – SMGPD SaJ 12, view from the right; E – SMGPD SaJ 24, view from the left, both from Hohnstein quarry (Germany);
F – NM-N96, F1 – lateral view, F2 – ventral view, Šternberk quarry; G – SMGPD SaJ 25, view from the right, Hohnstein quarry (Ger-
many); H – NM-N60, Kyjov. I—L – Vielunia sp.: I – CIGP 12651; J – NM-N172/1: J1 – view from the right, J2 – ventral view, Štern-
berk quarry; K – NM-N172/2; L – NM-N6, all from Šternberk quarry. M – Amoeboceras ovale Quenstedt, NM-N180: M1 – view from
the right, M2 – view from the left, Šternberk quarry. N – Amoeboceras (Plasmatites) praebauhini Salfeld, CIGP 10524, Šternberk quarry.
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density of secondaries. They have less prorsiradiate direc-
tion, sometimes almost radial, than primaries.
R e m a r k s : SMGPD SaJ 12: Rasenioides sp. figured here
resembles the Rasenia (Rasenioides) lepidula figured by
Geyer, 1961 (pl. 8, fig. 5, 6), but the characteristics of prima-
ries are worn away. SMGPD SaJ 24: Rasenia (Rasenioides)
paralepida (pl. 5, fig. 10) and R. (R.) transitoria (pl. 1,
fig. 6), both figured by Geyer, (1961) differ from this speci-
men figured by higher density of ribbing. A narrow umbili-
cal part with massive last whorl is typical. NM-N96:
Massive last whorl with poorly preserved umbilical part sug-
gests the affinity to the Rasenioides. SMGPD SaJ 25: Poorly
preserved mould similar to Rasenia (Rasenioides) transito-
ria figured by Geyer, 1961 (pl. 1, fig. 6). However, the fig-
ured specimen differs by having a more rursiradiate rib
direction. NM-N60: Massive outer whorl suggests the affinity
to Rasenioides.
Genus: Vielunia Wierzbowski & Glowniak, 2010
Vielunia sp.
Fig. 6I—L
M a t e r i a l : CIGP 12651: (fig. 6I), found in Šternberk
quarry near Brtníky. NM-N172/1: (fig. 6J), collected on pro-
file in Šternberk quarry near Krásná Lípa. NM-N172/2:
(fig. 6K), collected on profile in Šternberk quarry near Krásná
Lípa. NM-N6: (fig. 6L), found in Šternberk quarry near
Krásná Lípa.
Measurements: CIGP 12651: D (phg): 27 mm; Wh/D:
0.32; Wt/D: 0.35; U/D: 0.33. NM-N172/1: D (phg): 26 mm;
Wh/D: 0.37; Wt/D: 0.34; not measured. NM-N172/2:
D (phg-ad): 25 mm; Wh/D: 0.35; Wt/D: 0.32; not measured.
NM-N6: D (max): 29 mm.
D e s c r i p t i o n : Involute shells with a round or oval whorl
cross-section and a narrow umbilicus. Ribs have radial direc-
tion with a hint of falcoid rib type. Density of ribs is high
and they are thick and blunt.
R e m a r k s : NM-N172/1, 2: Only the last whorl shows
measurable parameters. Both specimens show involute
type of coiling with secondary filled umbilical area. The
density of ribbing on the last whorl is high. The figured
specimens resemble Vielunia dzalosinensis sp. nov. figured
by Wierzbowski et al., 2010 (pl. 9). NM-N6: Strongly de-
formed mould.
Genus: Aulacostephanoides
Aulacostephanoides sp.
Fig. 5K
M a t e r i a l : SMGPD SaJ 3: found in Hohnstein quarry,
Germany.
Measurements: D (max): 38 mm.
D e s c r i p t i o n : A part of massive whorl with high-square
whorl cross-section. Relatively short primary ribs show a
low density. Secondaries have higher density and fade on the
ventral margin, corresponding to the typical character of
Aulacostephanoides.
Genus: Aulacostephanus
Aulacostephanus sp.
Fig. 5L
M a t e r i a l : SMGPD SaJ 15: found in Hohnstein quarry,
(Germany).
Measurements: D (max): 70 mm.
D e s c r i p t i o n : A part of whorl with typically fading ribs
on the ventral margin, with a square-shaped whorl cross-sec-
tion and flattened lateral sides.
R e m a r k s : The weakening of the ribs suggests that this
specimen belongs to Aulacostephanus.
Family: Cardioceratidae Siemiradski, 1891
Genus: Amoeboceras Hyatt, 1900
Amoeboceras ovale Quenstedt, 1849
Fig. 6M
M a t e r i a l : NM-N180: Šternberk quarry.
Measurements: D (phg): 18 mm.
D e s c r i p t i o n : Involute type of shell. The figured speci-
men with partly filled umbilical part. The whorl cross-sec-
tion has ellipsoid shape. The density of ribbing is high. A
keel is developed on the ventral side, a typical character of
Amoeboceras. Relicts of sutures are recognizable on the lat-
eral side.
R e m a r k s : The specimen figured here shows a similarity
to Amoeboceras ovale figured by Gygi (2000), but the latter
has more flattened lateral sides. This specimen also resem-
bles A. ovale figured by Matyja & Wierzbowski, 1998 (pl. 1,
fig. 5, 6).
O c c u r r e n c e : Amoeboceras ovale occurs in the Regu-
lare and Rosenkrantzi Zones in the Boreal Realm and in the
upper part of the Bifurcatus Zone (Grossouvrei Subzone) up
to the lower part of the Bimammatum Zone (Hypselum Sub-
zone in the Sub-Mediterranean Realm).
Subgenus: Plasmatites Buckman, 1925
Amoeboceras (Plasmatites) praebauhini Salfeld, 1913
Fig. 6N
M a t e r i a l : CIGP 10524: Šternberk quarry.
Measurements: D (phg): 28 mm.
D e s c r i p t i o n : Involute coiling of shell, with secondarily
filled umbilical part. The specimen suggests an inversely
egg-shaped whorl cross-section. A typical keel is partly well
preserved.
R e m a r k s : Gently deformed and cracked carbonate
mould. Some ribs show variable direction caused by defor-
mation. The specimen figured shows a similarity with A.
praebauhini depicted by Atrops et al., 1993 (pl. 1, fig. 15—17).
A. bauhini figured by Schweigert & Callomon, 1997 (pl. 1)
suggests a similar deflection of ribs as the specimen figured
here.
Occurrence: Amoeboceras (Plasmatites) praebauhini oc-
curs in the uppermost part of the Bimammatum Zone (Hauf-
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fianum Subzone) and in the lower part of the Planula Zone in
the Sub-Mediterranean Realm. In the Sub-Boreal Realm,
taxon frequently occurs in the Baylei Zone (Densicostata and
Normandiana Subzones) as well as in the Bauhini and Kit-
chini Zones (Boreal Realm).
Geographical distribution and stratigraphic ranges
of Aulacostephanidae
Aulacostephanid ammonites appeared at the beginning of
the Late Oxfordian and became extinct at the end of the
Kimmeridgian (Page 2008). In addition to Sub-Boreal re-
gions, Aulacostephanidae were found in some Sub-Mediter-
ranean regions, such as southern Germany (Swabian Alb,
Franconian Alb), Swiss Jura and Central Poland (Schneid
1940; Dabrowska 1984; Gygi 2000, 2003; Matyja &
Wierzbowski 2002; Glowniak & Wierzbowski 2007; Hor-
nung 2009; Moliner 2009; Glowniak et al. 2010). Species of
this family were recorded also in the Boreal Realm, namely
in East Greenland, Norwegian Sea shelf, Spitsbergen, Franz
Josef Land, western and northern Siberia, British Columbia
and Canadian Arctic Archipelago (Mesezhnikov & Shulgina
1982; Mesezhnikov 1984, 1988; Birkelund & Callomon
1985; Wierzbowski 1989). However, cardioceratids are domi-
nant in the Boreal Realm and aulacostephanids are mostly
restricted to a few stratigraphic intervals (Rogov 2012). Aula-
costephanids are rare or entirely missing at most high-lati-
tude sites, for example, in northern Siberia (Mesezhnikov
1984; Rogov & Wierzbowski 2009; Wierzbowski & Rogov
2013).
Eight aulacostephanid genera (Microbiplices, Prorase-
nia, Rasenia, Eurasenia, Rasenioides, Vielunia, Aula-
costephanoides and Aulacostephanus) identified in the
Lower Kimmeridgian associations of northern Bohemia
represent new aulacostephanid occurrences in the Bohemian
Massif. The replacement of Microbiplices by younger
Eurasenia, Prorasenia or Rasenioides can correspond to
the Oxfordian/Kimmeridgian boundary interval in the Sub-
Boreal Realm (northern France, northern Germany and Great
Britain – Wright 2010). In contrast, the existence of these
aulacostephanid taxa indicates the Late Oxfordian in the
Sub-Mediterranean Realm (German-Polish branch). Aula-
costephanids appeared in Subpolar Urals during the latest
Oxfordian (Mesezhnikov 1984), and during the Early Kim-
meridgian became common in western and northern Siberia
(Rogov & Wierzbowski 2009).
Vielunia, first described by Wierzbowski (2010), occurs in
the upper part of the Bimammatum Zone and continues into
the Planula Zone, corresponding to the lowermost Kim-
meridgian in the Sub-Boreal and Boreal regions. The genus
Rasenia and related genera (i.e. Prorasenia, Eurasenia, and
Rasenioides) found in the Bohemian Massif belong to Sub-
Boreal ammonites. Rasenia has a relatively broad geographi-
cal range, covering Greenland, Spitsbergen, the Norwegian
Sea, Franz Josef Land, western Siberia, northern Siberia,
Great Britain, France, Germany, Poland, Russian Platform,
British Columbia and Arctic Canada (Tornquist 1896;
Schneid 1940; Birkelund et al. 1978, 1983; Callomon & Bir-
kelund 1980; Mesezhnikov 1984; Birkelund & Callomon
1985; Matyja et al. 2006; Wierzbowski et al. 2010). There-
fore, aulacostephanids from northern Bohemia and Saxony
thus seem to have a close relationship with ammonites repre-
senting the Sub-Mediterranean branch frequently occurring
in southern Germany and Poland.
Aulacostephanoides and Aulacostephanus belong to the
youngest genera of the family Aulacostephanidae described
here. Aulacostephanus was recorded in Sub-Boreal and Bo-
real Realm as well as in some Sub-Mediterranean regions. It
occurs in France, Great Britain, Switzerland, Germany, Nor-
way, East Greenland, Russian Platform, western and north-
ern Siberia (de Loriol 1874; Ziegler 1962; Birkelund et al.
1978, 1983; Mesezhnikov 1984; Callomon & Birkelund
1985).
Geographical distribution and stratigraphic ranges
of Cardioceratidae
The Cardioceratid genus Amoeboceras was described from
the Barents Sea, the Norwegian Sea shelf, Spitsbergen, west-
ern Siberia, northern Siberia, Russian Platform, and East
Greenland, (Sykes & Callomon 1979; Birkelund & Cal-
lomon 1985; Wierzbowski & Smelror 1993; Wierzbowski et
al. 2002; Meledina 2006; Rogov & Wierzbowski 2009;
Glowniak et al. 2010; Rogov 2010), as well as from Scot-
land, northern and central Poland, southern Germany, and
Switzerland (Matyja & Wierzbowski 1994, 2002; Schwei-
gert & Callomon 1997; Gygi 2000; Matyja et al. 2006;
Wierzbowski et al. 2010). The occurrence of Amoeboceras
in northern Bohemia corresponds to the equatorward migra-
tion of Boreal ammonites close to the Oxfordian/Kimmerid-
gian boundary. In the Polish Jura Chain and in southern
Germany, Amoeboceras (Plasmatites) praebauhini appears
in the upper part of the Sub-Mediterranean Bimammatum
Zone (Hauffianum Subzone) and continues into the Planula
Zone. This stratigraphic range correlates with the Early Kim-
meridgian Baylei and Bauhini Zones in Sub-Boreal and Bo-
real regions. A. ovale represents an older species occurring
during the Late Oxfordian and stratigraphically precedes
Amoeboceras (Plasmatites) praebauhini.
Discussion
Ammonite assemblages sharing many species in common
occur in the Polish Jura Chain, Holy Cross Mountains, Swa-
bian Alb and Franconian Alb (Wierzbowski 1978; Mali-
nowska 1991; Schweigert & Callomon 1997; Matyja &
Wierzbowski 2000, 2002; Wierzbowski et al. 2010). Ammo-
nites occur in deep neritic sponge megafacies in the Polish
Jura Chain as well as in southern Germany, and in more
shallow deposits of the carbonate platform of the Holy Cross
Mts. The connection between Poland and Germany that
crossed the Bohemian Massif is assumed to have been active
during the Late Jurassic (Matyja & Wierzbowski 1995,
2002; Schairer & Schlampp 2003). The connection between
these two areas enabled a migration of Boreal and Sub-Boreal
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ammonites across the Bohemian Massif southward to the
Tethyan Realm at the Oxfordian/Kimmeridgian transition
(Atrops et al. 1993; Matyja & Wierzbowski 1994, 2000).
Temporal proximity and lithological similarity of the Late
Jurassic sediments in the North Bohemian and Moravian ar-
eas (i.e. Brno vicinity, Olomučany village, etc.) evokes the
possibility of a paleogeographic connection between both re-
gions. However, the direct evidence is still missing because
no Jurassic sediments are preserved between these regions.
Conclusions
The Upper Jurassic marine deposits in northern Bohemian
Massif are rarely preserved and limited to a few small out-
crops, being mostly covered by Cretaceous and Tertiary de-
posits. The stretched fault system associated with the
Lusatian Fault, reactivated multiple times, does not allow a
discrimination of the original character of shelf and basinal
deposits. However, ammonites recovered from a few out-
crops and preserved in museum collections allow taxonomic
determinations and have implications for paleobiogeographic
pathways during the Late Jurassic. Aulacostephanid ammo-
nites suggest that the Late Jurassic carbonates belong to the
Upper Oxfordian and the Lower Kimmeridgian.
1. The appearance of typical Lower Kimmeridgian Boreal
and Sub-Boreal assemblages in northern Bohemia, repre-
sented by Prorasenia, Rasenioides or Eurasenia, marks the
Oxfordian/Kimmeridgian boundary.
2. With the exception of Prorasenia bathyschista Koerner
figured by Bruder in 1882, ammonites of family Aulaco-
stephanidae are described for the first time from the northern
Bohemian Massif.
3. The presence of Boreal and Sub-Boreal taxa in the
northern Bohemian Massif probably reflects the equatorward
migration of cold-water ammonites close to the Oxfordian/
Kimmeridgian boundary. Taxa occurring in northern Bohe-
mia show affinity to those occurring in the Polish Jura Chain
and southern Germany. This supports the presence of a ma-
rine connection between Poland and Germany across the Bo-
hemian Massif (Matyja & Wierzbowski (1995).
Acknowledgments: This paper was supported by Charles
University Grant Agency, Project No. 5947/2012, “Upper
Jurassic ammonites of northern Bohemia and their signifi-
cance”. The author is indebted to Dr. Jan Sklenář (National
Museum, Prague) for preparing excellent conditions during
my research in the National Museum. I also thank Dr.
Mikhail Rogov (GIN RAS Moscow), Dr. Andrzej
Wierzbowski (IG, University of Warsaw) and Dr. Günter
Schweigert (SMN, Stuttgart) for rigorous help with the de-
termination of ammonite specimens. Dr. Markus Wilmsen
allowed me access to the ammonite collections of the Na-
tional Museum in Dresden, Germany. I would like to thank
the Institute of Geology and Paleontology, Faculty of Sci-
ence, Charles University in Prague. I especially wish to
thank Dr. Martin Koš ák for very important and rewarding
consultation and supervision. Dr. Martin Mazuch and Dr.
Radek Vodrážka helped with access to fossil collections.
Finally, I would like to thank Dr. to Adam Tomašových for
valuable tutorial and final improvement of the English and
Mr. Alan Leath for English corrections.
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